Synopsis of Capparaceae to the flora of Colombia

A taxonomic synopsis of Capparaceae distributed in Colombia was carried out reviewing 1,800 botanical specimens from 13 herbaria in Colombia, five international institutions and close to 100 field collected specimens. We found 32 species and ten genera according to the last taxonomic modification proposed for this family. From these species, we reevaluated the conventional features used on the species identification, from which detailed description for each species and taxonomic keys were developed. In addition, geographic distribution maps in Colombia, and information about vernacular names and Neotropical distribution was added.

Antoine Laurent de Jussieu described Capparaceae in 1789 based on Capparis, from the Greek Kapparis or Kappari and Latin Capparis, which refers to shrubs or fruits of caper, Capparis spinosa L., (Quattrocchi 2000). After describing Capparis, Jussieu referred to the species of this group using the unofficial name Capparides. In this sense, the family name Capparidae was subsequently adopted in other taxonomic analyses (Bentham & Hooker 1862;De Candolle 1824) and, finally years later was named Capparidaceae (Bullock 1958(Bullock , 1959 and accepted by the International Rules of Botanical Nomenclature (Lanjow & Sprague 1947). However, in 1961 the Montreal code established that the correct name is Capparaceae based on current nomenclatural See supplementary material at <https: //doi.org/10.6084/m9.figshare.10141595.v1> Rodriguésia 70: e00232018. 2019 rules, e.g., dropping -is from Capparis and adding -aceae (International Association for Plant Taxonomy CIdlNB 1961). This modification was rejected by several authors who proposed keeping Capparidaceae because it came from the historical usage of Capparides, but the official name published by Jussieu was Capparis (Crosswhite & Iltis 1966;Dugand 1968b).
Nevertheless, Hutchinson (1967) in disagreement with the classification of Pax & Hoffmann (1936) proposed Capparidaceae with 32 genera and 440 species distributed in three subtribes (Apophylleae, Cadabeae, and Capparideae), without recognizing subfamilies. Hutchinson (1967) established this family in the order Capparales. Cronquist (1981) maintained Capparales but changed the name of the family to Capparaceae, divided it into the subfamilies Capparoideae and Cleomoidae, and recognized 45 genera and 800 species. Likewise, Takhtajan (1997) supported the classification of Cronquist (1981) but reduced the number of genera to 37 and increased the species to 900; he also insisted on adopting the name Capparaceae. Morton et al. (1997) removed the genus Physena to its own family, Physenaceae, and transferred it to Caryophyllales.
Subsequently, with molecular phylogenetic analyses, the classification system of Capparaceae has changed. A few analyses have found a close relationship between the subtribe Thelypodieae of Brassicaceae and Cleome of Cleomoideae (Judd et al. 1994). On the basis of these results Capparaceae was reduced to the level of subfamily and was included along with subfamily Cleomoideae in Brassicaceae s.l. Furthermore, other authors also suggested that Cleomoideae were more closely related with Brassicaceae than with Capparaceae (Rodman et al. 1993(Rodman et al. , 1996. These results led Hall et al. (2002) to analyze the phylogeny of Capparaceae and Brassicaceae using plastid genes, founding that Capparaceae s.l. was paraphyletic with respect to Brassicaceae.
However, evolutionary relationships within Capparaceae s.str. remained unclear, because the study by Hall et al. (2002) had limited taxon sampling. Hall (2008) performed another phylogenetic analysis of Capparaceae using two plastid genes and including 15 more species. The analysis clearly showed that Cleomoideae is more closely related to Brassicaceae with respect to Capparoideae, which led Hall (2008) to suggest a taxonomic revision separating the clades into three families, which were subsequently formally recognized by Iltis et al. (2011) as Capparaceae, Cleomaceae, and Brassicaceae. Another important outcome was that Capparis is paraphyletic, so it was proposed to segregate the genus into smaller clades for nomenclatural stability and monophyly. In this way species of Capparis were segregated into two large lineages, e.g. Capparis s.str. clade, which is predominantly Old World, and a clade of entirely New World Capparis, the latter more closely related to genera Atamisquea, Belencita, and Morisonia. Nonetheless, there are no useful morphological features for recognizing the New World taxa as a single genus. On the basis of these results, Iltis and Cornejo, in several publications, formalized the species of Neotropical capparoids as the following genera: Anisocapparis (1 species), Beautempsia (1) Calanthea (2), Caphexandrea (1), Capparicordis (3), Colicodendron (6), Capparidastrum (19), Cynophalla (16) Hispaniolanthus (1), Mesocapparis (1), Monilicarpa (2), Neocalyptrocalyx (7), Preslianthus (3), Quadrella (25), and Sarcotoxicum (1) (Cornejo et al. 2014;Cornejo 2010;Cornejo & Iltis 2005b, 2008b, c, e, 2009Galetti et al. 2016;Iltis 1965;Iltis & Cornejo 2007a, b, 2010a. There are no species of Capparis s.str. in the Neotropics. Over the last few years, several taxonomic monographs of this family have been published, mainly in Central America (Cornejo & Iltis 2012, 2013. In South America only new species (Cornejo & Iltis 2005a, 2010d, taxonomic combinations (Cornejo & Iltis 2006, 2008aIltis 2005;Rodríguez-Rodríguez et al. 2007) and regionals flora (Soares Neto et al. 2014;Soares Neto & Jardim 2015) have been published; there has not been modern monograph including South American species. In Colombia, Triana and Planchon (1862) recognized 21 species in Capparideae (Capparaceae), distributed in Cartagena, Santa Marta, and close to the Magdalena River. Subsequently, Dugand (1968aDugand ( , 1941 published a monograph of the genus Capparis in Colombia, as well as other studies that reported the genus Belencita (Dugand 1944 Iltis & Cornejo (2016) reported 29 species of Capparaceae in the catalogue of lichens and plants of Colombia (Bernal et al. 2016). However, the latter mainly included species deposited at the herbarium COL, and information about species distribution and identification keys is lacking. For this reason, the goal of this research is to carry out an extensive revision of specimens from all the herbaria of Colombia and several other major institutions in other countries, to make a detailed description, and an identification key focusing on vegetative characters; finally, we recorded the species distributions through the herbarium specimens exsiccata and stablished their geographic distribution.

Materials and Methods
To perform this study, we studied the nomenclatural types of the 29 species reported in Colombia, its taxonomic history, synonyms and valid name. With this information, we revised the protologues of each species and taxonomic types photography in JPS-Jstor Plant Science. Once we studied these taxonomic entities we reviewed the specimens distributed in Colombia at the national herbaria CAUP, COL, CUVC, FMB, HUA, HEUS, HECASA, ICESI, JAUM, MEDEL, UIS, UPTC, UTMC; and the foreign herbaria through online versions MO, K, NY, GH, and MEXU (all the herbarium acronym were carried out according to Holmgren et al. (1990) and updated based on Thiers (continuously updated).
After the herbarium specimens were reviewed, we used the geographic information from each specimen and elaborated distributions maps for each species of Capparaceae distributed in Colombia, employing Q-GIS ® (3.0). We used these dataset and maps to select locations that were mainly visited during 2015 and 2017 to collected plant specimens of Capparaceae.

Distributed in Venezuelan and Colombian
Andes, lowlands of Ecuador, Peru and Brazil. It is found in wet to very wet Andean forest and lowland forest (Fig. 1c). Cornejo & Iltis,Harvard Pap. Bot. 15 (1) Distributed from southwestern of Colombia to Equator. Found at wet to very wet Andean forest and lowland forest (Fig. 1c). Distributed from Mesoamerica to Caribbean region of Colombia. Found in tropical dry forest, disturbed forest and slightly wet forest (Fig. 1c). Distributed in the Colombian Amazonia, Venezuela and Brazil. Found in wet to very wet forest (Fig. 1c). Distributed from Mexico to Argentina. Found in a wet to very wet forest, lowland and tropical dry forest (Fig. 1d).